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ORIGIN OF THE METAZOANS 249
walled tube possibly of an unsegmented
worm. Distribution and ecology
Many of the Tommotian animals are form Although it is still unclear whether many of
taxa (that is, named simply by their shapes) the Tommotian skeletons are single shells or
because the biological relationships of most single sclerites and the autecology of most
cannot be established and often there are few groups is unknown, the assemblage was cer-
clues regarding the function and signifi cance tainly the first example in evolution of a skel-
of each skeletal part. Most are short-lived and etalized benthos. Very few of the Tommotian
have no obvious modern analogs. Two groups skeletal parts exceed 1 cm; nevertheless many
are common – the hyolithelminthids have shells were the armored parts of larger worm-
phosphatic tubes, open at both ends, whereas like animals. And both mobile and fi xed forms
the tommotiids are usually phosphatic, cone- occurred together with archaeocyathans and
shaped shells that seem to belong in bilater- non-articulate brachiopods. The microben-
ally symmetric sets. thos of the Tommotian was succeeded by a
Discoveries of near-complete examples of more typical Cambrian fauna, dominated by
Microdictyon-like animals from the Lower trilobites, non-articulate brachiopods, mono-
Cambrian of China have helped clarify the placophoran mollusks and primitive echino-
status and function of some elements of the derms together with the archaeocyathans
Tommotian fauna. These worms have round during the Atdabanian Stage (Fig. 10.14).
to oval plates arranged in pairs along the
length of the body, which may have provided
a base for muscle attachment associated with Cambrian explosion
locomotion. As noted previously, many of the The Cambrian explosion suddenly generated
small shelly fossils are probably the sclerites many entirely new and spectacular body plans
of larger multiplated worm and worm-like (Box 10.6) and coincides with the appearance
animals (Box 10.5). of the Bilateria over a relatively short period
of time (Conway Morris 1998, 2006). This
rapid diversification of life formed the basis
The Meishucunian biota for Stephen Jay Gould’s bestseller, Wonderful
Life (1989), which took its title from the
The Meishucunian Stage of South China has Frank Capra 1946 fi lm It’s a Wonderful Life.
yielded some of the most diverse Tommotian- The rapid appearance of such a wide range of
type assemblages in strata of Atdabanian age apparently different animals has suggested
(see Appendix 1). Qian Yi and Stefan Bengt- two possible explanations. The “standard”
son (1989) have described nearly 40 genera view is that the diversifi cation of bilaterians
that belong to three largely discrete, succes- happened just as fast as the fossils suggest,
sive assemblages through the stage. First, and that some reasons must be sought to
the Anabarites–Protohertzina–Arthrochites explain why many different animal groups
assemblage is dominated by tube-dwelling apparently acquired mineralized skeletons at
organisms such as Anabarites; the Siphongu- the same time. An alternative view arose after
chites–Paragloborilus assemblage contains initial molecular studies had suggested that
mobile mollusk-like and multiplated organ- animals diverged some 800 myr before the
isms together with some tube-dwellers and beginning of the Cambrian (e.g. Wray et al.
possible predators; whereas the Lapworth- 1996). If these molecular views were correct,
ella–Tannuolina–Sinosachites association has then the absence of fossils of modern animal
mainly widespread multiplated animals. phyla through the Proterozoic would have to
Many of these fossils are known from be explained by an interval of cryptic evolu-
Lower Cambrian horizons elsewhere in the tion of probable micro- and meioscopic organ-
world, highlighting the global distribution isms, living between grains of sand, operating
of many elements of the fauna. However, beneath the limits of detection prior to the
the three “community” types are rather explosion (Cooper & Fortey 1998). Greater
mysterious, and probably represent different refi nement of Cambrian stratigraphy, the tax-
ecosystems, but it is hard to speculate onomy and phylogeny of key Cambrian taxa
further. and their relative appearance in the fossil